Virtual reexamination of a plesiosaurian specimen (Reptilia, Plesiosauria) from the Late Cretaceous (Turonian) of Goulmima, Morocco, using computed tomography

LITERATURE CITED

• Andrews, C. W. 1909. On some new Plesiosauria from the Oxford Clay of Peterborough. Annals and Magazine of Natural History, London 48:418–429.
   Google Scholar
• Andrews, C. W. 1911. Description of a new plesiosaur (Plesiosaurus capensis, sp. nov.) from the Uitenhage Beds of Cape Colony. Annals of the South African Museum 1:309–322.
   Google Scholar
• Andrews, C. W. 1913. A Descriptive Catalogue of the Marine Reptiles of the Oxford Clay. Based on the Leeds Collection in British Museum (Natural History), Part II. British Museum (Natural History), London, 206 pp.
   Google Scholar
• Angst, D., and N. Bardet. 2015. A new record of the pliosaur Brachauchenius lucasi Williston, 1903 (Reptilia: Sauropterygia) of Turonian (Late Cretaceous) age, Morocco. Geological Magazine 153:449–459.
   Web of Science ®Google Scholar
• Araújo, R., M. J. Polcyn, J. Lindgren, L. L. Jacobs, A. S. Schulp, O. Mateus, A. O. Gonçalves, and M. L. Morais. 2015. New aristonectine elasmosaurid plesiosaur specimens from the Early Maastrichtian of Angola and comments on paedomorphism in plesiosaurs. Netherlands Journal of Geosciences-Geologie en Mijnbouw 94:93–108.
   Web of Science ®Google Scholar
• Bardet, N., P. Godefroit, and J. Sciau. 1999. A new elasmosaurid plesiosaur from the Lower Jurassic of Southern France. Palaeontology 42:927–952.
   Web of Science ®Google Scholar
• Bardet, N., X. Pereda Suberbiola, and N. E. Jalil. 2003a. A new polycotylid plesiosaur from the Late Cretaceous (Turonian) of Morocco. Comptes Rendus Palevol 2:307–315.
   Web of Science ®Google Scholar
• Bardet, N., X. Pereda Suberbiola, and N. E. Jalil. 2003b. A new mosasauroid (Squamata) from the Late Cretaceous (Turonian) of Morocco. Comptes Rendus Palevol 2:607–616.
   Web of Science ®Google Scholar
• Bardet, N., A. Houssaye, J. C. Rage, and X. Pereda Suberbiola. 2008. The Cenomanian–Turonian (Late Cretaceous) radiation of marine squamates (Reptilia): the role of the Mediterranean Tethys. Bulletin de la Société Géologique de France 179:605–622.
   Web of Science ®Google Scholar
• Benson, R. B. J., and P. S. Druckenmiller. 2014. Faunal turnover of marine tetrapods during the Jurassic–Cretaceous transition. Biological Reviews 89:1–23.
   PubMed Web of Science ®Google Scholar
• Benson, R. B. J., M. Evans, and P. S. Druckenmiller. 2012. High diversity, low disparity and small body size in plesiosaurs (Reptilia, Sauropterygia) from the Triassic–Jurassic boundary. PLoS ONE 7:e31838. doi: 10.1371/journal.pone.0031838
   PubMed Web of Science ®Google Scholar
• Benson, R. B. J., K. T. Bates, M. R. Johnson, and P. J. Withers. 2011. Cranial anatomy of Thalassiodracon hawkinsii (Reptilia, Plesiosauria) from the Early Jurassic of Somerset, United Kingdom. Journal of Vertebrate Paleontology 31:562–574.
   Web of Science ®Google Scholar
• Blainville, H. D. de 1835. Description de quelques espèces de reptiles de la Californie, précédée de l'analyse d'un système général d'Erpétologie et d'Amphibiologie. Nouvelles Annales du Muséum (national) d'Histoire Naturelle de Paris 4:233–296.
   Google Scholar
• Brown, D. S. 1993. A taxonomic reappraisal of the families Elasmosauridae and Cryptoclididae (Reptilia: Plesiosauria). Revue de Paléobiologie 7:9–16.
   Google Scholar
• Brown, D. S., P. Vincent, and N. Bardet. 2013. Osteological redescription of the skull of Microcleidus homalospondylus (Sauropterygia, Plesiosauria) from the Lower Jurassic of England. Journal of Paleontology 87:537–549.
   Web of Science ®Google Scholar
• Buchy, M. C. 2005. An elasmosaur (Reptilia: Sauropterygia) from the Turonian (Upper Cretaceous) of Morocco. Carolinea 63:5–28.
   Google Scholar
• Buchy, M. C., F. Métayer, and E. Frey. 2005. Osteology of Manemergus anguirostris n. gen. et sp., a new plesiosaur (Reptilia, Sauropterygia) from the Upper Cretaceous of Morocco. Palaeontographica, Abteilung A 272:97–120.
   Google Scholar
• Cabrera, A. 1941. Un plesiosaurio nuevo del Cretáceo del Chubut. Revisita del Museo de la Plata (Neuvo Serio) 2:113–130.
   Google Scholar
• Carpenter, K. 1997. Comparative cranial anatomy of two North American Cretaceous plesiosaurs; pp. 191–216 in J. M. Callaway and E. L. Nicholls (eds.), Ancient Marine Reptiles. Academic Press, San Diego, California.
   Google Scholar
• Carpenter, K. 1999. Revision of North American elasmosaurs from the Cretaceous of the Western Interior. Paludicola 2:148–173.
   Google Scholar
• Cavin, L. 1995. Goulmimichthys arambourgi n. g., n. sp., un Pachyrhizodontidae (Actinopterygii, Teleostei) d'une nouvelle localité a nodules fossilifères du Turonien inferieur marocain. Comptes Rendus de l'Académie des Sciences, Paris, Série IIa 321:1049–1054.
   Google Scholar
• Cavin, L. 1997. Nouveaux Teleostei du gisement du Turonien inférieur de Goulmima (Maroc). Compte Rendus de l'Académie des Sciences de Paris 325:19–24.
   Google Scholar
• Cavin, L. 1999. Occurrence of a juvenile teleost, Enchodus sp., in a fish gut content from the Upper Cretaceous of Goulmima, Morocco. Palaeontology 60:57–72.
   Google Scholar
• Cavin, L. 2001. Osteology and phylogenetic relationships of the teleost Goulmimichthys arambourgi Cavin, 1995 from the Upper Cretaceous of Goulmima, Morocco. Eclogae Geologicae Helvetica 133:25–52.
   Google Scholar
• Cavin, L., L. Boudad, S. Duffaud, L. Kabiri, J. Le Loeuff, I. Rouget, and H. Tong. 2001. L’évolution paléoenvironnementale des faunes de poissons du Crétacé supérieur du bassin du Tafilalt et des régions avoisinantes (Sud-Est du Maroc): implications paléobiogéographiques. Compte Rendu de l'Académie des Sciences de Paris, Sciences de la Terre et des planètes 333:677–683.
   Google Scholar
• Cavin, L., H. Tong, L. Boudad, C. Meister, A. Piuz, J. Tabouelle, M. Aarab, R. Amiot, E. Buffetaut, G. Dyke, S. Hua, and J. Le Loeuff. 2010. Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: an overview. Journal of African Earth Sciences 57:391–412.
   Web of Science ®Google Scholar
• Cheng, Y. N., T. Sato, X. C. Wu, and C. Li. 2006. First complete pistosaurid from the Triassic of China. Journal of Vertebrate Paleontology 26:501–504.
   Web of Science ®Google Scholar
• Cope, E. D. 1869. Synopsis of the extinct Batrachia and Reptilia of North America. Transactions of the North American Philosophical Society 14:1–252.
   Google Scholar
• Cruickshank, A. R. I. 1997. A Lower Cretaceous pliosauroid from South Africa. Annals of the South African Museum 105:207–226.
   Google Scholar
• Cruickshank, A. R., and R. E. Fordyce. 2002. A new marine reptile (Sauropterygia) from New Zealand: further evidence for a Late Cretaceous austral radiation of cryptoclidid plesiosaurs. Palaeontology 45:557–575.
   Web of Science ®Google Scholar
• Dames, W. 1895. Die Plesiosaurier der süddeutschen Liasformation. Abhandlungen der Königliche Preussische Akademie der Wissenschaften zu Berlin 1895:1–83.
   Google Scholar
• Druckenmiller, P. S. 2002. Osteology of a new plesiosaur from the Lower Cretaceous (Albian) Thermopolis Shale of Montana. Journal of Vertebrate Paleontology 22:29–42.
   Web of Science ®Google Scholar
• Druckenmiller, P. S., and A. P. Russell. 2008. A phylogeny of Plesiosauria (Sauropterygia) and its bearing on the systematic status of Leptocleidus Andrews, 1922. Zootaxa 1863:1–120.
   Google Scholar
• Ettachfini, E. M., and B. Andreu. 2004. Le Cénomanien et le Turonien de la plate-forme Préafricaine du Maroc. Cretaceous Research 25:277–302.
   Web of Science ®Google Scholar
• Evans, M. 2012. A new genus of plesiosaur (Reptilia: Sauropterygia) from the Pliensbachian (Early Jurassic) of England, and a phylogeny of the Plesiosauria. Ph.D. dissertation, University of Leicester, Leicester, U.K., 397 pp.
   Google Scholar
• Ferrandini, M., J. Philip, J. F. Babinot, and G. Tronchetti. 1985. La plate-forme carbonatée du Cénomano–Turonien de la région d'Erfoud-Errechidia (Sud-Est marocain): stratigraphie et paléoenvironement. Bulletin de la Société Géologique de France 8:559–564.
   Google Scholar
• Gasparini, Z., L. Salgado, and S. Casadi. 2003a. Maastrichtian plesiosaurs from northern Patagonia. Cretaceous Research 14:157–170.
   Web of Science ®Google Scholar
• Gasparini, Z., N. Bardet, J. E. Martin, and M. Fernandez. 2003b. The elasmosaurid plesiosaur Aristonectes Cabrera from the latest Cretaceous of South America and Antarctica. Journal of Vertebrate Paleontology 23:104–115.
   Web of Science ®Google Scholar
• Goloboff, P. A., J. M. Carpenter, J. S. Arias, and D. R. M. Esquivel. 2008. Weighting against homoplasy improves phylogenetic analysis of morphological datasets. Cladistics 24:1–16.
   Web of Science ®Google Scholar
• Großmann, F. 2007. Preliminary description and phylogenetic position of a new plesiosaur (Reptilia: Sauropterygia) from the Toarcian of Holzmaden, Germany. Journal of Paleontology 78:973–988.
   Google Scholar
• Kear, B. P. 2005. A new elasmosaurid plesiosaur from the Lower Cretaceous of Queensland, Australia. Journal of Vertebrate Paleontology 25:792–805.
   Web of Science ®Google Scholar
• Ketchum, H. F., and R. B. J. Benson. 2010. Global interrelationships of Plesiosauria (Reptilia, Sauropterygia) and the pivotal role of taxon sampling in determining the outcome of phylogenetic analyses. Biological Reviews 85:361–392.
   PubMed Web of Science ®Google Scholar
• Kubo, T., M. T. Mitchell, and D. M. Henderson. 2012. Albertonectes vanderveldei, a new elasmosaur (Reptilia, Sauropterygia) from the Upper Cretaceous of Alberta. Journal of Vertebrate Paleontology 32:557–572.
   Web of Science ®Google Scholar
• Lebedel, V., C. Lezin, B. Andreu, M. J. Wallez, E. M. Ettachfini, and L. Riquier. 2013. Geochemical and palaeoecological record of the Cenomanian–Turonian Anoxic Event in the carbonate platform of the Preafrican Trough, Morocco. Palaeogeography, Palaeoclimatology, Palaeoecology 369:79–98.
   Web of Science ®Google Scholar
• Lomax, D. R., and W. R. Wahl. 2013. A new specimen of the elasmosaurid plesiosaur Zarafasaura oceanis from the Upper Cretaceous (Maastrichtian) of Morocco. Paludicola 9:97–109.
   Google Scholar
• Maisch, M. 1998. Notes on the cranial osteology of Muraenosaurus Seeley, 1874 (Sauropterygia, Jurassic), with special reference to the neurocranium and its implications for sauropterygian phylogeny. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 207:207–253.
   Google Scholar
• Novas, F. E., J. S. D'Angelo, J. P. O'Gorman, F. L. Agnolín, J. M. Lirio, and M. P. Isasi. 2015. First record of Polycotylidae (Sauropterygia, plesiosauria) from the Upper Cretaceous of Antarctica. Cretaceous Research 56:563–568.
   Web of Science ®Google Scholar
• O'Gorman, J. P., L. Salgado, E. B. Olivero, and S. A. Marenssi. 2015. Vegasaurus molyi, gen. et sp. nov. (Plesiosauria, Elasmosauridae), from the Cape Lamb Member (lower Maastrichtian) of the Snow Hill Island Formation, Vega Island, Antarctica, and remarks on Wedellian Elasmosauridae. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2014.931285
   Web of Science ®Google Scholar
• O'Gorman, J. P., R. A. Otero, N. Hiller, J. Simes, and M. Terezow. 2017. Redescription of Tuarangisaurus keyesi (Sauropterygia; Elasmosauridae), a key species from the Uppermost Cretaceous of the Weddellian province: internal skull anatomy and phylogenetic position. Cretaceous Research 71:118–136.
   Web of Science ®Google Scholar
• O'Keefe, R. F. 2001. A cladistic analysis and taxonomic revision of the Plesiosauria (Reptilia: Sauropterygia). Acta Zoologica Fennica 213:1–63.
   Google Scholar
• O'Keefe, R. F. 2006. Morphologic and ontogenetic patterns in elasmosaur neck length, with comments on the taxonomic utility of neck length variables. Paludicola 5:207–229.
   Google Scholar
• Otero, R. A., J. P. O'Gorman, N. Hiller, F. R. O'Keefe, and R. E. Fordyce. 2016. Alexandronectes zealandiensis gen. et sp. nov., a new aristonectine plesiosaur from the lower Maastrichtian of New Zealand. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2015.1054494
   PubMed Web of Science ®Google Scholar
• Otero, R. A., S. Soto-Acuña, F. R. O'Keefe, J. P. O'Gorman, W. Stinnesbeck, M. E. Suárez, D. Rubilar-Rogers, C. Salazar, and L. A. Quinzio-Sinn. 2014. Aristonectes quiriquinensis, sp. nov., a new highly derived elasmosaurid from the upper Maastrichtian of central Chile. Journal of Vertebrate Paleontology 34:100–125.
   Web of Science ®Google Scholar
• Owen, R. 1860. On the orders of fossil and Recent Reptilia, and their distribution through time. Report of the British Association for the Advancement of Science 1859:153–166.
   Google Scholar
• Sachs, S. 2005. Redescription of Elasmosaurus platyurus Cope 1868 (Plesiosauria: Elasmosauridae) from the Upper Cretaceous (lower Campanian) of Kansas, U.S.A. Paludicola 5:92–106.
   Google Scholar
• Sachs, S., and B. P. Kear. 2015. Postcranium of the paradigm elasmosaurid plesiosaurian Libonectes morgani (Welles, 1949). Geological Magazine 152:694–710.
   Web of Science ®Google Scholar
• Sachs, S., and B. P. Kear. 2017. Redescription of the elasmosaurid plesiosaurians Libonectes atlasense from the Upper Cretaceous of Morocco. Cretaceous Research 74:205–222.
   Google Scholar
• Sachs, S., J. Lindgren, and M. Siversson. 2015. A partial plesiosaurian braincase from the Upper Cretaceous of Sweden. Geological Society, London, Special Publications 434:293–301.
   Google Scholar
• Sato, T. 2002. Description of plesiosaurs (Reptilia: Sauropterygia) from the Bearpaw Formation (Campanian–Maastrichtian) and a phylogenetic analysis of the Elasmosauridae. Ph.D. dissertation, University of Calgary, Calgary, Alberta, Canada, 391 pp.
   Google Scholar
• Sato, T. 2003. Terminonatator ponteixensis, a new elasmosaur (Reptilia: Sauropterygia) from the Upper Cretaceous of Saskatchewan. Journal of Vertebrate Paleontology 23:89–103.
   Web of Science ®Google Scholar
• Sato, T., Y. Hasegawa, and M. Manabe. 2006. A new elasmosaurid plesiosaur from the Upper Cretaceous of Fukushima, Japan. Palaeontology 49:467–484.
   Web of Science ®Google Scholar
• Sato, T., X. C. Wu, A. Tirabasso, and P. Bloskie. 2011. Braincase of a polycotylid plesiosaur (Reptilia: Sauropterygia) from the Upper Cretaceous of Manitoba, Canada. Journal of Vertebrate Paleontology 31:313–329.
   Web of Science ®Google Scholar
• Smith, A. S., and G. J. Dyke. 2008. The skull of the giant predatory pliosaur Rhomaleosaurus cramptoni: implications for plesiosaur phylogenetics. Naturwissenschaften 95:975–980.
   PubMed Web of Science ®Google Scholar
• Storrs, G. W., and M. A. Taylor. 1996. Cranial anatomy of a new plesiosaur genus from the lowermost Lias (Rhaetian/Hettangian) of Street, Somerset, England. Journal of Vertebrate Paleontology 16:403–420.
   Web of Science ®Google Scholar
• Stromer, E. 1935. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wusten Ägyptens. Wirbeltierreste der Baharijestufe (unterstes Cenoman), 15. Plesiosauria: Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse 26:1–56.
   Google Scholar
• Taylor, M. A. 1992. Functional anatomy of the head of the large aquatic predator Rhomaleosaurus zetlandicus (Plesiosauria; Reptilia) from the Toarcian (Lower Jurassic) of Yorkshire, England. Philosophical Transactions of the Royal Society of London, Series B 335:247–280.
   Web of Science ®Google Scholar
• Underwood, C., K. M. Cleason, and D. J. Ward. 2009. Batoids from the Turonian of SE Morocco; p. 43 in N. E. Jalil (ed.), 1st International Congress on North African Vertebrate Palaeontology, Program and Abstracts, Marrakech, 25–27 May 2009.
   Google Scholar
• Vincent, P., N. Bardet, A. Houssaye, M. Amaghzaz, and S. Meslouh. 2013. New plesiosaur specimens from the Maastrichtian Phosphates of Morocco and their implications for the ecology of the latest Cretaceous marine apex predators. Gondwana Research 24:796–805.
   Web of Science ®Google Scholar
• Vincent, P., N. Bardet, X. Pereda Suberbiola, B. Bouya, M. Amaghzaz, and S. Meslouh. 2011. Zarafasaura oceanis, a new elasmosaurid (Reptilia: Sauropterygia) from the Maastrichtian Phosphates of Morocco and the palaeobiogeography of latest Cretaceous plesiosaurs. Gondwana Research 19:1062–1073.
   Web of Science ®Google Scholar
• Welles, S. P. 1943. Elasmosaurid plesiosaurs with a description of new material from California. Memoirs of the University of California 13:125–254.
   Google Scholar
• Welles, S. P. 1949. A new elasmosaur from the Eagle Ford Shale of Texas. Fordren Science, Series 1:1–28.
   Google Scholar
• Welles, S. P. 1952. A review of the North American Cretaceous elasmosaurs. University of California, Publications in Geological Sciences 29:47–144.
   Google Scholar
• Welles, S. P. 1962. A new species of elasmosaur from the Aptian of Columbia and a review of the Cretaceous plesiosaurs. University of California Publications in Geological Sciences 44:1–96.
   Google Scholar
• Wiffen, J., and W. L. Moisley. 1986. Late Cretaceous reptiles (families Elasmosauridae and Pliosauridae) from the Mangahouga Stream, North Island, New Zealand. Journal of Geology and Geophysics, New Zealand 29:205–252.
   Web of Science ®Google Scholar
• Williston, S. W. 1889. A new plesiosaur from the Niobrara Cretaceous of Kansas. Transactions of the Annual Meetings of the Kansas Academy of Science 12:174–178.
   Google Scholar
• Williston, S. W. 1903. North American plesiosaurs. Part I. Field Columbian Museum Publication 2:1–79.
   Google Scholar
• Handling editor: Patrick Druckenmiller.
   Google Scholar